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Hydrodynamically controlled anagenetic of from Poland

MARIUSZ NIECHWEDOWICZ and JERZY TRAMMER

Niechwedowicz, M. and Trammer, J. 2007. Hydrodynamically controlled anagenetic evolution of Famennian goniatites from Poland. Acta Palaeontologica Polonica 52 (1): 63–75.

This paper reports on the evolution of ammonoids belonging to the family from the of Janczyce in the Holy Cross Mountains, Poland. Steady and gradual changes in conch morphology of the lin− eage frechi– subacutum–T. sublentiforme occurred in concert with water shallowing during the deposition of the Lower Famennian . Biometric analysis of ammonoid conch and facies analysis of the cephalopod have been applied to assess the possible relationship between shell geometry and envi− ronmental changes. Results show that ratios of width / diameter as well as whorl width / whorl height decreased, while distance from the venter to the greatest whorl width / diameter increased with time, thereby reducing hydrody− namic drag of the shells, probably in response to increasing water turbulence. The interpretation presented here is in agreement with similar cases from the literature. However, this kind of environmentally controlled evolution has hith− erto been recognized only in and ammonoids. Conch morphology may be considered as an indica− tor of palaeobathymetry.

Key words: , hydrodynamics, evolution, environmental changes, Devonian, Poland.

Mariusz Niechwedowicz [[email protected]], Muzeum Wydziału Geologii, Uniwersytet Warszawski, ul. Żwirki i Wigury 93, PL−02−089 Warszawa, Poland; Jerzy Trammer [[email protected]], Instytut Geologii Podstawowej, Wydział Geologii, Uniwersytet Warszawski, ul. Żwirki i Wigury 93, PL−02−089 Warszawa, Poland.

Introdution The extraordinarily rich material from the lower Famen− nian cephalopod limestone from Janczyce (Holy Cross Ammonoids are one of the most prodigious groups of Mountains, Poland) offers new insights into patterns and ; they have long attracted students of evolution due to factors controlling ammonoid evolution in the Palaeozoic. their outstanding fossil record, extraordinary diversification, The assemblage from Janczyce is strongly dominated by wide distribution and rapid . While their fossil re− tornoceratid goniatitids, a group with worldwide distribu− cord is well documented, patterns of ammonoid evolution tion. Its representatives are common in the Middle and Up− have yet to be fully explored. The most commonly postulated per Devonian rocks of (Holy Cross Mountains, factor controlling ammonoid evolution is sea level−fluctu− Rhenish Slate Mountains, Harz, Montagne Noire, Pyre− ation (e.g., Bayer and McGhee 1984; House 1993; Korn nees, Cantabrian Mountains), North Africa (Morocco, Al− 1995; Klug 2002; Wani 2003), which may lead to paedo− geria), Asia (Ural, Timan), Australia (Canning Basin), and morphosis, as documented by Korn (1995) for the Late De− (e.g., House and Price 1985; Becker 1993; vonian ammonoids, as well as to other evolutionary changes Becker et al. 2000). in ammonoid shell morphology. Commonly, such patterns of Institutional abbreviation.—MWG, Museum of the Faculty ammonoid shell modification are observed in successions re− of Geology, Warsaw University, Warsaw, Poland. cording the regressive phases of transgressive−regressive cy− cles. Falls in relative sea−level and increases in water turbu− lence can trigger evolutionary changes manifested in gradual modifiactions in ammonoid shell geometry. This pheno− Geological setting menon has been well documented among ammo− noids (e.g., Bayer and McGhee 1984; Jacobs et al. 1994). The goniatitid assemblage described herein was collected However, it is poorly known in their Palaeozoic counter− from the Janczyce section (GPS position data: 50°46’1.20” N, parts. This paucity of data on evolutionary patterns among 21°13’58.80” E), in the eastern Holy Cross Mountains, Po− goniatitids is apparently caused by the scarcity of appropriate land (Fig. 1). The outcrop is situated in the eastern part of the sections for population studies of this group. Łagów Syncline in the Kielce Region. The goniatitids were

Acta Palaeontol. Pol. 52 (1): 63–75, 2007 http://app.pan.pl/acta52/app52−063.pdf 64 ACTA PALAEONTOLOGICA POLONICA 52 (1), 2007

Fig. 1. Location of the Janczyce area (A) within the Holy Cross Mountains (B) and its position within Poland (C). The Janczyce outcrop is marked with an asterisk. Map of the Holy Cross Mountains modified after Mizerski (1995). Janczyce geological map after Narkiewicz and Olkowicz−Paprocka (1983). collected from a 1.2 metre thick cephalopod limestone, part The cephalopod limestone of the Janczyce section was of the over 200 metres thick Łagów Beds (Makowski 1991; found in 1918 by Jan Samsonowicz, who mapped this area Matyja and Narkiewicz 1995; see Fig. 2). This unit is a series geologically (Makowski 1991). On the basis of goniatitid of generally poorly fossiliferous (except for small brachio− material from this section, Makowski (1962, 1991) described pods, tentaculitids, and rare goniatitids), regulary bedded, . dark−grey laminated marly limestones (Narkiewicz and Ol− According to Makowski (1991), the Janczyce cephalopod kowicz−Paprocka 1983). limestone is lenticular. It splits into a number of layers which The Janczyce cephalopod limestone contains a rich cepha− differ in colour and faunal composition. Makowski (1991) lopod fauna (goniatitids, cyrto− and orthoconic ) to− recognized five beds within this limestone (Fig. 2). He also gether with rare brachiopods, gastropods, and bivalves. These noted the presence of two sedimentary discontinuity surfaces rocks are very similar in lithology and age to other cephalopod stained with oxidized iron minerals. These mark the bound− limestone intercalations known from Europe and other parts of aries between Beds 1–2 and 3–4. the world. They occur in the Upper Devonian of the Rhenish The age of the cephalopod limestone has been determined Slate Mountains and the Harz Mountains (Germany), the using conodonts. Wolska (1967) identified the Fammenian Montagne Noire (France), the Moravian Karst (Czech Repub− crepida Zone. More detailed studies have fur− lic), the Carnic (Austria, Italy), the Cantabrian Moun− ther constrained the age to Middle (Szulczewski 1992; Matyja tains (Spain), and in the Anti−Atlas chains of southern Mo− and Narkiewicz 1995) or Upper Palmatolepis crepida Zone rocco (Wendt and Aigner 1985). (according to Woroncowa−Marcinowska 2002). NIECHWEDOWICZ AND TRAMMER—ANAGENETIC EVOLUTION OF GONIATITES 65 Material and methods Bed 5 The material used in this study was collected from a temporary outcrop—a trench dug in a field of a local farmer. After the fieldwork was finished, the exposure was covered. It is be− lieved that the Janczyce site may be completely worked−out. During the fieldwork in 2001 and 2002 a few hundred tornoceratid goniatites were collected. The first rich ammo− noid material coming from this locality was collected by Bed 4 Henryk Makowski in the middle of the twentieth century. Makowski was interested in the problem of sexual dimor− phism in ammonites, so that he collected and described only sedimentary adult specimens. Moreover, he gave only the diameters of his discontinuity specimens and did not measure other parameters of the shell (see Makowski 1991). We also noticed the presence of juve− nile and adolescent specimens. Therefore, both collections Bed 3 were compared, but only the present authors’ collection was Palmatolepis crepida used in this study. The measurements were taken from all ontogenetic stages of the ammonoids found. Accordingly, the original variability spectrum of the “palaeopopulations” for each part of the log was reproduced as accurately as pos− Bed 2 sible. FAMENNIAN sedimentary Ammonoid specimens, especially large ones, are usually 1m incompletely preserved. The body chamber is usually dam− Middle / Upper Zone discontinuity aged, and fragments of the are preserved only in a few specimens. However, most of the goniatitids extracted 0 from the rocks are three−dimensionally preserved, which al− Bed 1 lowed us to use them in this study. The body chambers are marly limestones usually filled with silty sediment while the shales 10 cm consist of clear calcite or are filled with silty sediment, which cephalopod limestones caused damage of the septa. Therefore in most specimens the early whorls cannot be observed. Ammonoids with incom− Fig. 2. Position of the studied Janczyce cephalopod limestones within the plete conchs, usually caused by body chamber damage, spec− complex of laminated marly limestones (Łagów Beds). Cephalopod lime− imen corrosion or erosion, were not used in this work. Many stone log after Makowski (1991). specimens have the wrinkle−layer structure preserved (see Niechwedowicz 2003) which is interpreted as “fingerprint”− junior synonym of T. typum by Korn and Klug (2002). How− like impression of the soft body (e.g., Senior 1971; Dogu− ever, differences in conch morphology and line be− zhaeva and Mutvei 1996). tween these forms allow us to regard Tornoceras sublenti− forme as a valid taxon. Becker (1993) suggested possible relationships between Description of goniatitid some representatives of the studied Tornoceratidae lineage and other Late Devonian species. According to him, T. sub− morphotypes acutum and T. sublentiforme are closely related to the oxy− conic Oxytornoceras acutum. However, there are some dif− Most of the goniatitids examined belong to the family Torno− ferences between these forms, especially in the suture line ceratidae. The remainder represent the family Cheilocera− and shell ornamentation (presence of furrows in O. acutum; tidae, including subpartitum (Münster, 1839) see Becker 1993: 363). Moreover, O. acutum is considerably which is the index fossil of the Cheiloceras II a Zone (zona− smaller and stratigraphically younger. Thus, T. subacutum tion of Wedekind 1913). and T. sublentiforme seem to represent endemic forms that Makowski (1991) was the first to observe the evolution− evolved in isolation, probably only in the Holy Cross Moun− ary lineage within Tornoceratidae from the Janczyce section. tains Basin. Representatives of this lineage are also known This evolutionary lineage is composed of three major spe− from other localities in the Holy Cross Mountains: Kadziel− cies: Phoenixites frechi (Wedekind, 1918), Tornoceras sub− nia (Makowski 1991) and Jabłonna (Jerzy Dzik, personal acutum Makowski, 1991 and Tornoceras sublentiforme So− communication 2003); see Fig. 1. These taxa probably evol− bolew, 1914 (see Fig. 3). T. sublentiforme was regarded as a ved from Phoenixites frechi, which is a long−ranging and

http://app.pan.pl/acta52/app52−063.pdf 66 ACTA PALAEONTOLOGICA POLONICA 52 (1), 2007

III

Bed 5

Bed 4

II

Bed 3

Bed 2 I

Bed 1

. m 0.1

Fig. 3. Typical representatives of morphotype populations occurring in the cephalopod limestone sequence in Janczyce, Holy Cross Mountains, Poland. The log has been split into three parts, each of them containing different ammonoid assemblages. I (Beds 1, 2) contain Phoenixites frechi (Wedekind, 1918) specimens: MWG ZI/08/006/25 (A), MWG ZI/36/015 (B), MWG ZI/08/067 (C), and MWG ZI/08/024 (D); note also the presence of other goniatitids: Falcitornoceras aff. korni (Becker, 1984), MWG ZI/08/283 (E), Simicheiloceras cf. schrickeli Becker, 1993, MWG ZI/36/294 (F), and Cheiloceras (Cheiloceras) subpartitum (Münster, 1839), MWG ZI/36/293 (G). II (Bed 3) contains Tornoceras subacutum Makowski, 1991 specimens: MWG ZI/36/152 (H), MWG ZI/08/040 (I), and MWG ZI/08/042 (J), some of ammonoids belonging to this group (those with rounded venter) display intermediate characters between Phoenixites frechi–Tornoceras subacutum. III (Beds 4, 5) contain Tornoceras sublentiforme Sobolew, 1914 specimens: MWG ZI/08/008/54 (K), MWG ZI/08/055 (L), and MWG ZI/08/009/17 (M). A1–K1, lateral view; A2–K2, ventral view. Note the trend in morphological shell modifications observed on ascending the log. Scale bars 10 mm. NIECHWEDOWICZ AND TRAMMER—ANAGENETIC EVOLUTION OF GONIATITES 67

20 20 III 10 10 Bed 5

10 10 5 5 N=54

37.1 0.321 0.563 0.457

20 40 60 0.28 0.36 0.44 0.52 0.5 0.7 0.9 0.36 0.44 0.52 Bed 4

20 20 20 20 II N=71 10 10 10 10 Bed 3

31.8 0.347 0.603 0.384

20 40 60 0.28 0.36 0.44 0.52 0.5 0.7 0.9 0.36 0.44 0.52

Number of specimens I Bed 2 50 30 40 20 20 N=131 30 20 Bed 1 20 10 10 10 10 28.9 0.4 0.695 0.366

20 40 60 mm 0.28 0.36 0.44 0.52 0.5 0.7 0.9 0.36 0.44 0.52 ww f dm Tr = ww S= F= dm wh2 dm

statistical significance (p = 0.01) wh1 statistical significance (p = 0.05) dm mean statistical significance (p = 0.001) 0.347 wh2 f minimal and maximal value mean confidence interval (p = 0.95) standart deviation ww

Fig. 4. Results of the biometrical analyses. Trends in shell shape modifications can be clearly seen in succeeding beds of the analysed section. The values of the morphological parameters Tr and S decrease with time while the values of the parameter F tend to increase upwards through the succession. This is ex− pressed in the disc−shaped and more compressed conchs of the ammonoids from Beds 4 and 5. Mean values of the diameter also increase upwards. Note that differences between the means of the morphological parameters (dm, Tr, S, and F) are statistically significant (p = 0.05–0.001). See the text for the details. I–III, morphotype populations. geographically widespread Devonian form (Becker 1993; phology of those ammonoids differs considerably, their early Becker et al. 2000). ontogenetic stages are nearly identical. Hence all specimens Each species belonging to the studied lineage occurs in a were assigned to one species. By analogy, it cannot be ex− different part of the succession (see Fig. 3) and exhibits high cluded that all ammonoids from Janczyce cephalopod lime− intraspecific variation (see histograms in Fig. 4). Some of the stone might also belong to a single species. We had difficul− specimens are intermediate forms between named taxa and ties in identifying some specimens, especially from the group cannot be easily assigned to any of them. This confirms that II in Fig. 3, as some of them (e.g., Fig. 3J) show intermediate the succession of goniatitids found in the Janczyce section is characters between Phoenixites frechi–Tornoceras subacu− a good example of an anagenetic evolutionary lineage. tum. It clearly indicates that the systematics of the Fam− Another example shows a different possible interpreta− menian goniatites requires thorough revision. This is, how− tion of within ammonoids. An ammonoid assem− ever, beyond the scope of our paper. blage from the of Siberia, with similar variability in Because of the various ways in which the fauna studied shell morphology to those from Janczyce, was studied by may be interpreted taxonomically, the goniatitids were in− Dagys and Weitschat (1993). Although adult conch mor− vestigated not as species but rather as populations of mor−

http://app.pan.pl/acta52/app52−063.pdf 68 ACTA PALAEONTOLOGICA POLONICA 52 (1), 2007 photypes from successive Beds. They were separated into (see Fig. 4): Tr, thickness ratio—whorl width (ww)/diame− the following three morphotype populations (Fig. 3): ter (dm), S, shape of aperture (= relative whorl thick− (I) Morphotypes with involute, slightly depressed disco− ness)—whorl width (ww)/ whorl height (wh2), and F,flank conic shells, with a rounded venter; found in Beds 1 and 2; position—distance from the venter to the greatest whorl see Fig. 3 (I). Type−representative taxon of this group is width (f)/diameter (dm). Phoenixites frechi. This sample consists of 131 specimens. Two of the Raupian coiling parameters (D, distance from (II) Morphotypes with a more compressed, involute the coiling axis and W, whorl expansion rate; see Raup 1967) conch and a rounded to subtrapezoidal shaped venter, found were not used because all of the ammonoids investigated have in Bed 3. 71 specimens were used in the study, with Torno− involute conchs (with nearly the same D values) and have the ceras subacutum as a typical representative; see Fig. 3 (II). same whorl expansion rate (W). Moreover, thickness ratio (Tr) (III) The most slender morphotypes among those investi− and whorl shape (S) influence drag to a greater extent than the gated; have involute, oxyconic shells, with a sharp−edged coiling parameters D and W and thus are of greater relevance venter, some forms having a slightly rounded venter and in in ammonoid hydrodynamics (Jacobs 1992). some a slight keel appears but only on the body chamber; found in Beds 4 and 5; see Fig. 3 (III). Tornoceras sublenti− forme is typical of this group. The sample consists of 54 specimens. Results All studied Tornoceratidae have involute conchs. The body chamber extends more than 270°. Suture lines are Thickness ratio (Tr) and shape of aperture (S).—These pa− very similar in all populations (see Niechwedowicz 2003). rameters describe the slenderness of the conch (Tr) and the The slenderness of the shell in almost all specimens studied whorl (S). Their values tend to decrease with time. In the up− can be categorized as either less compressed (Tr = 0.3–0.5; per Beds (4 and 5) the most slender ammonoids occur. The sensu Jacobs 1992) or compressed with Tr mean values less greatest mean value changes in Tr and S occur between the than 0.3 (for definition of morphological parameters see populations from Beds 1–2 and Bed 3 (Tr mean value chan− Fig. 4). ges from 0.400 to 0.347, and S mean value from 0.695 to Besides the goniatite species mentioned above, the Jan− 0.603; see Fig. 4). All observed differences between mean czyce cephalopod limestone also contains other tornoceratids: values are statistically significant (p = 0.001). Falcitornoceras aff. korni (Becker, 1984), Simicheiloceras cf. Flank position (F).—This parameter describes the position schrickeli Becker, 1993 (see Fig. 3), and Exotornoceras su− of the widest cross−section of the whorl. Its values increases perstes (Wedekind, 1908). upwards through the succession. The greatest values of whorl width for ammonoids from the lower beds occur close to the venter, while in ammonoids from the upper beds they Biometric analysis lie close to . The greatest modifications of F mean values are—in contrast to Tr and S parameters—between Measurements of the ammonoid conch were taken from Bed 3 and Beds 4–5 (F mean values change from 0.384 to the body chamber (as shown in Fig. 4) using calipers and 0.457; see Fig. 4). This is expressed in the disc−shaped conch rounded to 0.1 mm. Altogether, 256 specimens have been of the ammonoids from Beds 4 and 5. Differences between measured: 131 from the first morphotype population (Beds mean values of this parameter are statistically significant (p = 1–2), 71 from the second (Bed 3) and 54 specimens from the 0.001). third (Beds 4–5). As mentioned above, the measurements were all taken from ammonoid growth stages present within Diameter (dm).—Mean values of diameter also increase with the Janczyce cephalopod limestone. Sexual dimorphism, al− time and differences between means are significant (Fig. 4). though suggested by Makowski (1962, 1991), appears to be This biometrical analysis clearly shows how representa− doubtful within the studied assemblage and was not taken tives of the investigated ammonoid population became into account in this investigation. Results of measurements more compressed and disc−shaped with time. Particularly are shown in Fig. 4. notable is that the evolution of this lineage progressed in The simplest way to document evolutionary trends is to easily identifiable stages. The greatest changes in conch plot morphological parameters through time. Raupian coil− slenderness (as expressed in Tr and S values) are observed ing parameters (see Raup 1967) are useful for describing between Beds 1–2 and Bed 3 (see Fig. 4). This may be con− ammonoid spiral shell geometries, under the assumption of sidered as the first stage of shell shape modification (“thick− isometric growth, and have been used by numerous ammo− ness stage”). In contrast, the coiling parameter F shows the nite workers (e.g., Ward 1980; Bayer and McGhee 1984; greatest modification in mean value between Bed 3 and Jacobs 1992; Korn 2000; Klug 2001; Korn and Klug 2001, Beds 4–5, while Tr and S both tend to change more slowly. 2003). These parameters as well as the F coiling parameter, Moreover, during this stage the ventral part of goniatitid proposedbyChamberlain(1976),werealsousedinthis shells tends to become sharper. This is the second stage of study. The morphological variables applied in this paper are morphological shell modification. NIECHWEDOWICZ AND TRAMMER—ANAGENETIC EVOLUTION OF GONIATITES 69

1994). This could be a result of reduced mechanical strength Hydromechanical design of the of oxyconic whorls. goniatitid shells from Janczyce Accordingly, habitat depth limitation can be calculated on the basis of theoretical modelling of hydrodynamic drag It is generally accepted that most ammonoids were poor swim− and of implosion depth of ammonoid shells. Tornoceras mers (e.g., Kennedy and Cobban 1976; Lewy 2002). They uniangulare, which is very similar in shell morphology and were not well designed for continuous rapid swimming and sutural complexity to Phoenixites frechi from Beds 1 and 2 of acceleration (Jacobs and Chamberlain 1996). Some ammo− the Janczyce section, yielded a depth estimate of 160 metres noid workers (e.g., Chamberlain 1976; Bayer and McGhee (Hewitt 1996). The oxyconic ammonoid Beloceras, similar 1984; Jacobs 1992; Jacobs et al. 1994) have suggested that in shell geometry to Tornoceras sublentiforme (occurring in there is a close relationship between mode of life and shell Beds 4–5), has a much shallower habitat depth limit, calcu− morphology in ammonoids. According to these authors, the lated at 53 metres (Hewitt 1996). ammonoid shell produced hydrodynamic drag during swim− Thus, shells of more depressed ammonoids were presum− ming. Shell shape, as mathematically defined, allows hydro− ably poorer in swimming under high−energy water flow condi− dynamic efficiency and ammonoid swimming ability to be tions but were resistant to hydrostatic pressure. Their power calculated. The great disparity in ammonoid conch morpho− consumption level was optimal at greater depths, while com− logies suggests that some ammonoids swam slowly, others pressed oxyconic ammonoids supposedly preferred much more rapidly. shallower environments because of lower tolerance of hydro− Experiments show that water movement near the shell sur− static pressure and higher hydrodynamic efficiency. Depres− face during ammonoid swimming was not the same on each sed forms were possibly well adapted to slow, continuous part of conch. The greatest water turbulence was created in the swimming but moderately or poorly adapted to fast continu− umbilicus and behind the shell (Chamberlain 1976). Moreover, ous swimming and acceleration. This is the opposite to com− the greater the area of ammonoid cross−section the greater the pressed, oxyconic forms which seem to have been good at fast drag that is produced. On the other hand, slender shells cannot continuous swimming and acceleration, but rather moderate accommodate as much muscular tissue for swimming as de− or even poor at slow, continuous swimming (Jacobs and pressed shells. Therefore, compressed ammonoids might have Chamberlain 1996). achieved lower hydrodynamic efficiency (Kazushige Tanabe, Moreover, the size of ammonoid shells influences drag and personal communication 2006). Muscle scars are not preserved power consumption during swimming (Jacobs and Cham− in the investigated material and therefore hydrodynamic effi− berlain 1996). With increasing size, drag increases as a func− ciency cannot be calculated in this way. tion of area whereas power availability increases as a function Ammonoids would have been most hydrodynamically ef− of volume (Jacobs 1992). Additionally, larger forms travelling ficient in fast swimming when their shells were as slender as at higher velocities are more efficient if they are more com− possible, with little or no umbilicus and with a sharp−edged pressed (lower value of Tr). This can be concluded from the venter. In biometric terms this kind of shell will have low Tr distribution of conch forms in the Janczyce succession (see and S but high F values. Oxyconic shell shapes match this de− Fig. 4). scription very well. Increases in the prominence of shell ornamentation would This is not, however, the ideal type of ammonoid conch. If also increase hydrodynamic drag. In the Janczyce material this it had been, all ammonoids should have been oxyconic. Ex− factor is insignificant because all ammonoids have smooth periments show that compressed ammonoids were most hy− conchs. drodynamically efficient only in high−energy environments (e.g., Jacobs et al. 1994). This means that oxycones could not swim fast but rather were able to cope with high−energy water flow. In deeper, usually low−energy environments, this shell Facies analysis of the cephalopod morphology was not preferred as the energy consumption of limestones oxyconic ammonoids was apparently much greater than de− pressed ammonoids (Jacobs 1992). Facies analysis has been applied in examining the depo− There is one additional force (other than drag) which oper− sitional environment of the Janczyce cephalopod limesto− ates in water environments. This is hydrostatic pressure, the nes and their possible correlations with trends in ammonoid value of which increases with depth. Compilation of am− shell shape. The following facies characteristics were ob− monoid sutural complexity and shape of whorl cross−section served: allows calculation of shell strength and resistance against the force of hydrostatic pressure. Implosion depth of the shell and Beds 1–2.—(1) Dark grey in colour; (2) containing more de− likely habitat depth limit can be calculated (Hewitt 1996). pressed goniatitids with a rounded venter: Phoenixites frechi, Hewitt (1996) observed that habitat depth limit of depressed Cheiloceras (Cheiloceras) subpartitum, Simicheiloceras cf. ammonoids was much deeper, even by comparison with more schrickeli, Falcitornoceras aff. korni, and Exotornoceras compressed varieties of the same taxon (see also Jacobs et al. superstes, Fig. 3 (I); (3) cyrto− and orthoconic nautiloids cha−

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g

e n

c jg

g o

jg

Fig. 5. Thin sections from the Janczyce section. A. Laminated marly limestone (MWG ZI//36/296); sampled below the cephalopod limestone log. B. Cephalopod wackestone (MWG ZI//36/301); Bed 1. Note the presence of goniatitids (g), orthoconic nautiloids (n), gastropods (ga), entomozoacean ostracodes (e) and poor sorting of bioclasts. C. Cephalopod wackestone (MWG ZI//36/303); Bed 3. Note the presence of goniatitids (g), juvenile goniatitids (jg), crinoids (c) and molluscan shell debris. D. Cephalopod packstone (MWG ZI//36/305); Bed 5. Note the presence of juvenile goniatitids (jg), benthic ostracode (o). Note also the best sorting of bioclasts in this bed. Scale bars 2 mm. otically dispersed; (4) placoderm remains; (5) pelagic ostra− only compressed, oxyconic forms occur (Tornoceras sublenti− cods (family Entomozoidae, Fig. 5B); (6) maximum abun− forme, Fig. 3 (III)); (3) absence of cyrtoconic nautiloids; only dance of crinoids and gastropods; (7) unsorted, small to large orthoconic forms occur, usually oriented (Fig. 6); (4) benthic bioclasts. ostracodes (Fig. 5D); (5) crinoids and gastropods rare or ab− sent; (6) well−sorted bioclasts, usually smaller in comparison Bed 3.—(1) Black in colour; (2) containing more com− with those from Beds 1–3; much less matrix. pressed goniatitids with a subtrapezoidal venter (Tornoceras subacutum), some forms with a slightly rounded venter (tran− sitional forms: Phoenixites frechi–Tornoceras subacutum, Fig. 3 (II); (3) very few cyrtoconic nautiloids; (4) moderate Depositional environment abundance of crinoids and gastropods. A number of workers (e.g., Tucker 1973; Wendt et al. 1984; Beds 4–5.—(1) Black (Bed 4) or grey (Bed 5) in colour; (2) Wendt and Aigner 1985; Santantonio 1994; Szulczewski et

Fig. 6. A. Current− or wave−oriented orthoconic nautiloids (MWG ZI/08/285); Bed 5 of the Janczyce cephalopod limestone. B. Current− or wave−oriented ® orthoconic nautiloids (MWG ZI/08/284) occurring together with Tornoceras sublentiforme (Ts); Bed 5 of the Janczyce cephalopod limestone. Arrows indi− cate current or wave direction. Scale bar 10 mm. NIECHWEDOWICZ AND TRAMMER—ANAGENETIC EVOLUTION OF GONIATITES 71

Ts

Ts

Ts

Ts

http://app.pan.pl/acta52/app52−063.pdf 72 ACTA PALAEONTOLOGICA POLONICA 52 (1), 2007 al. 1996) have expressed the opinion that cephalopod lime− 1–2. All these goniatitids have a more depressed conch with stones were often deposited on pelagic carbonate platforms a rounded venter, like Phoenixites frechi which occurs in the and platform slopes. These platforms were usually tectonic same beds. or volcanic in origin. Because of strong stratigraphic conden− (3) Upwards decrease in the abundance of cyrtoconic sation and sedimentary discontinuity surfaces, sedimentary nautiloids, with no specimens found in Beds 4 and 5. rate must have been very low (1 to 5 m per million ; (4) Chaotic orientation of orthoconic nautiloids in the Wendt and Aigner 1985). The depositional depth of cephalo− lower layers of the sequence as compared with current− or pod limestones has been calculated at several tens to several wave−orientation in Beds 4–5; this suggests shallower, even hundreds of metres (Wendt and Aigner 1985). subtidal, high−energy environments (e.g., Wendt and Aigner The depositional environment of the Janczyce cephalopod 1985; Wendt 1995). limestone seems to have been very similar to other examples (5) Changes in the ostracod fauna, with deep−water ento− of this type of rock. During the Middle and Late Devonian the mozoids (see Olempska 1992) being found only in the lower palaeogeography of the Holy Cross Mountains was dominated part of sequence, while benthic ostracodes occurred in the by a shallow−marine carbonate platform (Szulczewski 1995). upper Beds. Its growth caused the separation of northern and southern (6) Placoderm remains have been noted only from the intra−shelf basins. During the late and the early Fa− lower part of the Janczyce section. mennian the platform was drowned and transformed into a pe− (7) Upward decrease in crinoids and gastropods. lagic carbonate platform. The northern margin of the shal− (8) Upward decrease in . In the upper layers, low−marine carbonate platform, where Janczyce area was situ− only one species of goniatitid (Tornoceras sublentiforme) ated, drowned in the Late Frasnian, marking the beginning of occurs and the benthic fauna is clearly impoverished. monotonous, laminated marl basin sedimentation in this area. (9) Upward decrease in abundance of large bioclasts and The pelagic carbonate platform was characterised by differen− matrix, and increase in fine shell debris, suggesting a high− tial bottom morphology, probably caused by Late Devonian energy environment for Beds 4–5 (microfacies changing extensional tectonics as suggested, among others, by Racki from wackestones in Beds 1–2 to packstones in the upper and Narkiewicz (2000). This resulted in the formation of iso− beds). lated carbonate platforms succeeding the condensed cephalo− The microfacies in the lower part of the cephalopod lime− pod limestones (Szulczewski 1995; Szulczewski et al. 1996). stone shows some differences in comparison with Beds 4 and The origin of pelagic carbonate platforms is connected to 5. However, these probably are not significant (Stanisław early−orogenic block faulting and differential subsidence (simi− Skompski, personal communication 2002). Tucker (1973) lar trends were reported from Morocco by Wendt et al. 1984). suggested that diagenetic processes could have obliterated Little is known about the palaeogeography of the eastern the original texture of the Devonian cephalopod limestones. part of the Holy Cross Mountains because there are very few Thus, other facies features seem to be more useful as palaeo− outcrops. However, some sedimentary structures have been environmental indicators. described from this area. Slump structures indicating north− Macrofacies analysis clearly shows that the upper beds of wards transportation of sediment were noted from Łagów the Janczyce cephalopod limestone were deposited in a higher (Radwański and Roniewicz 1962) (see Fig. 1), similar in age energy regime than the lower layers. This high−energy envi− to the Janczyce cephalopod limestone, may be evidence of ronment was probably a result of bottom current or wave ac− the existence of a tectonically active isolated pelagic carbon− tivity, as indicated by parallel alignment of orthoconic nauti− ate platform in the Janczyce area. The cephalopod limestone loids. The Janczyce cephalopod limestone sequence seems to must have been deposited on the top of this pelagic platform. represent a shallowing−upward cycle. Sedimentary discontinuities described by Makowski (1991) Another possibility—post mortem shell accumulation— as well as the great abundance of conodonts (Tatiana Woron− that could account for the origin of these ammonoid−bearing cowa−Marcinowska, personal communication 2002) suggest sediments can be excluded. Most of the goniatitids from the condensation. This is in agreement with the cephalopod Janczyce cephalopod limestone are preserved with the body limestone depositional model. chamber intact. This suggests that the ammonoid shells were Trends in faunal and facies changes discovered in the not transported very far to their site of deposition (e.g., Korn Janczyce sequence may be summarised as follows: 1995). The presence of juvenile goniatitid shells (Fig. 5) that (1) The earliest tornoceratid morphotypes in the succes− do not show extensive breakage also suggests that the gonia− sion were depressed forms with a rounded venter (Beds 1–2). titids were deposited within the habitat realm of the popula− These were followed by more compressed forms with a tion (see Tanabe et al. 1993). Such a great number of am− subtrapezoidal venter (Bed 3) and finally, the most com− monoid specimens in the Janczyce cephalopod limestone pressed, oxyconic forms with a sharp−edged venter (Beds 4 seems to be the result of condensation, which has often been and 5). noted for cephalopod limestones. The presence of sedimen− (2) Other goniatitid genera occur only in the lower part of tary discontinuity surfaces noted by Makowski (1991) sup− the sequence: specimens of Cheiloceras, Simicheiloceras, ports this interpretation. The large number of conodont spec− Falcitornoceras, and Exotornoceras were found in Beds imens also favours stratigraphic condensation. NIECHWEDOWICZ AND TRAMMER—ANAGENETIC EVOLUTION OF GONIATITES 73

The conclusion here is that the two factors of block−tecton− Discussion ics and eustatic sea−level fluctuations must have influenced fa− cies distribution in the Holy Cross Mountains during the Late Ammonoid abundance relates closely to sea−level fluctua− Devonian (e.g., Szulczewski 1989). A relative sea−level fall tions, being generally greater when sea−level is high and less during the Middle/Late Palmatolepis crepida Zone may have when it is low (e.g., House 1993). This is the result of reduc− also happened in the Janczyce area. This is consistent with our tion in habitat area during regression which may lead to ex− proposed cephalopod limestone facies interpretation. tinction, known as the species−area effect (Wani 2003). The Hydromechanical analyses of the ammonoids studied to− opposite can happen as well, as documented by Monnet et al. gether with the facies analysis allow calculation of approxi− (2003). mate depositional depths. Depositional depth of the lower To survive environmental changes, ammonoids had to layers of Janczyce cephalopod limestone can be estimated at adapt (Bayer and McGhee 1984; Jacobs et al. 1994; Korn about 160 metres using the formula of Hewitt (1996). Similar 1995; Seki et al. 2000; Kawabe 2003). According to these neritic depth values were suggested for the deposition of authors, sea−level change is a very important factor promot− laminated marly limestones (Flügel 1982) like those associ− ing ammonoid evolution; they suggest that ammonoids were ated with the Janczyce cephalopod limestone. In contrast, the restricted to some extent to environments hydrodynamically upper beds of the cephalopod limestone sequence may have appropriate to their shell morphologies. Compressed, oxy− been deposited at only a few metres depth. conic ammonoids were better adapted for living in high−en− A cephalopod limestone intercalation of similar age in the ergy, usually shallow−water environments, while depressed Kadzielnia Quarry is also thought to have been deposited forms probably had optimal power consumption in low−en− during the same regressive phase (Narkiewicz 1987). Al− ergy, generally deeper environments. though this outcrop is located a few tens of kilometres west The likelihood of relative sea−level changes in the Holy of Janczyce (see Fig. 1) and is situated on the southern mar− Cross Mountains during the Middle/Late Palmatolepis cre− gin of the Holy Cross Mountains Famennian pelagic carbon− pida Zone time has been suggested by other authors. Neptu− ate platform, there are similarities between the localities. nean dykes noted from the Kadzielnia Quarry in Kielce are Makowski (1991) noted the presence of a very similar torno− the result of an extensional tectonic regime (Racki and Nar− ceratid lineage within the Kadzielnia cephalopod limestone. kiewicz 2000). This kind of regime leads to block−faulting Cephalopod limestones from these localities differ in colour, and thus to varying bottom palaeomorphology. Tectonic microfacies, and associated facies. In addition, the Kadziel− block uplifts might have led to local shallowing or even nia cephalopod limestone contains very few . How− emergence of the block, as has been described for Gałęzice ever, the presence of a very similar ammonoid succession (Szulczewski et al. 1996). suggests similar depositional environments. Eustatic sea−level changes might also have occurred. Narkiewicz (1987) postulated a local regression during the Middle Palmatolepis crepida Zone. This may be correlated with a small regressive pulse just before the eustatic trans− Conclusions gression described by Johnson et al. (1985). However, the occurrence of this event in the Holy Cross Mountains is This paper reports on the morphological evolution of Famen− doubted by some authors (Szulczewski 1990). Also during nian Tornoceratidae from the Holy Cross Mountains (Po− the Late Palmatolepis crepida Zone, a global regressive land). Biometrical analysis of goniatitids, together with fa− pulse has been suggested by some authors (Dopieralska cies analysis of the cephalopod limestone, leads to the fol− 2003; Schülke and Popp 2005). Dopieralska (2003) used lowing conclusions: neodymium (Nd) isotopic composition of conodonts as a (1) Biometrical analysis shows various trends in shell tool to investigate Late Devonian sea−level fluctuations. shape changes in this ammonoid lineage. These are de− She concluded that the Nd isotopic composition of cono− creases in the values of thickness ratio (Tr) and shape of the donts reflects sea−level fluctuations, with values decreasing aperture (S), and increase in the values of flank position (F) during the regressive phases. According to Dopieralska, the coiling parameters; regressive event during the Late Palmatolepis crepida Zone (2) The parameters Tr, S,andF appear to have the greatest was one of five prominent eustatic events during the late relevance to ammonoid hydrodynamics. As with analogous Frasnian and early Famennian (see also Buggisch 1991). examples from the literature, these trends in the values of coil− Apart from this event, the two Kellwasser events, also re− ing parameters are interpreted as adaptations in shell shape corded as condensed cephalopod facies, were deposited among ammonoids living in high−energy environments. This during regressive pulses (Dopieralska 2003). These results, interpretation is supported by the trend in morphology of the however, contrast with the opinion of some other workers venter geometry: from round through trapezoidal to sharp− (e.g., Szulczewski 1995; Racki and Narkiewicz 2000) who edged. Oxyconic ammonoids, representing the final stage in have suggested that cephalopod limestone deposition is as− shell shape, occur in the upper part of the Janczyce sequence. sociated with periods of deepening. This kind of conch morphology is thought to have the lowest

http://app.pan.pl/acta52/app52−063.pdf 74 ACTA PALAEONTOLOGICA POLONICA 52 (1), 2007 power consumption during swimming in high−energy envi− Buggisch, W. 1991. The global Frasnian–Famennian “Kellwasser Event”. ronments; International Journal of Earth Sciences 80: 49–72. Chamberlain, J.A. Jr. 1976. Flow pattern and drag coefficients of cephalopod (3) Facies analysis of the cephalopod limestone suggests a shells. Palaeontology 19: 539–563. shallowing−upwards cycle. The presence of pelagic ostra− Dagys, A.S. and Weitschat, W. 1993. Extensive intraspecific variation in a codes (family Entomozoidae) in the lower part of the succes− Triassic ammonoid from Siberia. Lethaia 26: 113–121. sion, benthic shallow−water ostracodes and current− or wave− Doguzhaeva, L. and Mutvei, H. 1996. Attachment of the body to the shell in oriented orthoconic nautiloids in the upper part and general ammonoids. In: N.H. Landman, K. Tanabe, and R.A. Davis (eds.), upward biodiversity decrease confirms this inference; Ammonoid Paleobiology. Topics in Geobiology 13: 44–64. Dopieralska, J. 2003. Neodymium Isotopic Composition of Conodonts as a (4) Relative sea−level fall, proposed in this case, may Palaeoceanographic Proxy in the Variscan Oceanic System. Disserta− have been caused by tectonic uplift and/or eustatic regres− tion Universität Giessen. 111 pp. Giessener Elektronische Bibliothek. sion. Both factors may have been involved, since extensional URL:http://geb.uni−giessen.de/geb/volltexte/2003/1168/; URN:urn:nbn: tectonic movements and eustatic regressive pulses both oc− de: hebis: 26−opus−11682 curred in the Middle/Late Palmatolepis crepida Zone when Flügel, E. 1982. Microfacies Analysis of Limestones. 633 pp. Springer−Verlag, Berlin. the Janczyce cephalopod limestone was deposited; Hewitt, R. A. 1996. Architecture and strength of the ammonoid shell. In: (5) Morphological evolution of the studied tornoceratid N.H. Landman, K. Tanabe, and R.A. Davis (eds.), Ammonoid Paleo− lineage appears to be associated with a relative sea−level fall. biology. Topics in Geobiology 13: 297–344. This provides one more example of this kind of environmen− House, M.R. 1993. Fluctuations in ammonoid evolution and possible envi− tally controlled evolution, but is probably the first docu− ronmental controls. In: M.R. House (ed.), The Ammonoidea, Environ− mented in Palaeozoic ammonoids. This can be considered as ment, Ecology, and Evolutionary Change. Systematics Association Spe− cial Volume 47: 13–34. a subsidiary tool for interpreting the relative depth of sedi− House, M.R. and Price, J.D. 1985. New Late Devonian genera and species ment deposition. However, much more data from other lin− of tornoceratid goniatites. Palaeontology 28: 159–188. eages and ages is needed to support a relationship between Jacobs, D.K. 1992. Shape, drag, and power in ammonoid swimming. Paleo− sea−level change and evolutionary trends in ammonoid shell. biology 18: 203–220. Jacobs, D.K. and Chamberlain, J.A. Jr. 1996. Buoyancy and hydrodynamics in ammonoids. In: N.H. Landman, K. Tanabe, and R.A. Davis (eds.), Ammonoid Paleobiology. Topics in Geobiology 13: 169–224. Acknowledgements Jacobs, D.K., Landmann, N.H., and Chamberlain, J.A. Jr. 1994. Ammonite shell shape covaries with facies and hydrodynamics: iterative evolution as a response to changes in basinal environment. Geology 22: 905–908. We are grateful to Ewa Olempska−Roniewicz (Institute of Paleobio− Johnson, J.G., Klapper, G., and Sandberg, C.A. 1985. Devonian eustatic logy, Polish Academy of Sciences, Warsaw, Poland) and Stanisław fluctuation in Euramerica. Geological Society of America Bulletin 96: Skompski (Faculty of Geology, Warsaw University, Poland) for con− 567–587. structive discussion on microfacies analysis. We also thank Maciej Kawabe, F. 2003. 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